Pseudophallus starksii

Dallevo-Gomes, Caio I. A., Mattox, George M. T. & Toledo-Piza, Mônica, 2020, Taxonomic review of the pipefish genus Pseudophallus Herald, with the description of a new species (Syngnathiformes: Syngnathidae), Zootaxa 4859 (1), pp. 81-112 : 98-102

Pseudophallus starksii ( Jordan & Culver 1895)

( Figs. 2–3 , 5 , 7 ; Table 2)

Siphostoma starksii Jordan & Culver (in Jordan, 1895): 381, 416, 514, pl. XXX (original description; “Río Presidio, about a mile below from village Presidio, Sinaloa, Mexico ”; type specimens SU 2686); Jordan & Evermann 1896b: 328 (listed in North America); Jordan & Evermann, 1898: 2838 (redescription; type listed as SU 2686); Jordan & Evermann, 1900: 3259, pl. CXX:3 25 (explanation of pl. CXX, drawing of SU 2686); Böhlke, 1953: 62 (syntypes listed); Dawson, 1982: 47 (synonymic list of P. starksii ).

Siphostoma starksi (sic): Jordan & Evermann, 1896a: 771 (, redescription, type listed as USNM 47425); Rutter, 1896: 263 (occurrence in San José del Cabo, Baja California, Mexico); Meek, 1904: 162–163 (occurrence in Río San Geronimo, Mexico, key of identification).

Siphostoma (Siphostoma) starksii : Jordan & Evermann, 1896b: 328, 576 (new combination; listed in Central America); Dawson, 1982: 47 (synonymic list of P. starksii ).

Syngnathus starksi (sic): Starks, 1906: 783 (new combination, occurrence in Santa Rosa, Ecuador); Regan, 1907: 55 (listed); Jordan et al., 1930: 242 (listed in Central America); De Buen, 1940: 46 (listed in Mexico).

Siphostoma elcapitanense : (non Meek & Hildebrand, 1914): Meek & Hildebrand (in Meek, 1914): 119 (misidentification; Jesús Maria and Turrubales, Costa Rica); Dawson, 1982: 47 (synonymic list of P. starksii ).

Syngnathus elcapitanense : (non Meek & Hildebrand, 1914): Meek & Hildebrand, 1923: 260 (in part: incorrect identification of specimens from Costa Rica and Punta Chame, Panama); Dawson, 1982: 47 (synonymic list of P. starksii ).

Pseudophallus starksii: Gunter, 1942: 310 (euryhaline); Fowler, 1944: 496 (listed in Central America); Gunter, 1956: 349 (listed in Central America); Follett, 1960: 213, 214, 224 (mention of affinities with congeners); Dawson, 1982: 45–49, 165, figs. 17, 20, 70 (species account, comparisons with congeners, identification key; specimen CAS(SU) 67185 (ex SU 2686) as lectotype); Dawson, 1985: 161, 162, 209, figs. 240, 241 (map) (characters, distribution, distribution, comparisons with other species of Pseudophallus , identification key); Ferraris Jr., 2003: 591 (listed, distribution; SU 67185 (ex SU 2686) as lectotype); Ruiz-Campos et al., 2002: 148, 149 (listed in Baja California Sur, Mexico); Nelson et al., 2004: 112 (listed for Central America); Robles & Vega, 2004: 82 (listed in Santa Maria river, Veraguas, Panama); Kuiter, 2009: 241 (distribution, freshwater, figure from Dawson, 1982); Mol, 2012: 51 (citation of literature regarding positive rheotropism); McMahan et al., 2013: 448 (listed in El Salvador); Page et al., 2013: 110 (listed in Central America); González-Acosta et al., 2015: 72 (listed in Baja California, Mexico); Angulo et al., 2015: 7, 9, 10; fig. 5K (listed, distribution in Costa Rica, identification key); Angulo et al., 2020: 9; fig. 8N (as starski), 19, 28 (as starksii , listed in Costa Rica, identification key).

Pseudophallus starksi (sic): Herald, 1940: 51, 59, 63 (new combination; designated as type species of Pseudophallus ; species account; identification key); Herald 1959: 473 (general occurrence in the Pacific coast); Miller, 1966: 796 (coastal streams, San José del Cabo to Ecuador); Bussing, 1966: 222, 236 (occurrence in Costa Rica. listed); Whatley, 1969: 448 (cited as euryhaline fish); Dawson, 1974:409, 410 (meristic characters in tables and identification key); Lagler et al., 1977: 39 (figure from Jordan & Evermann 1896); Villano & Balderas, 1987: 230 (listed in Mexico); Castro-Aguirre et al., 1995: 72 (listed in Gulf of California, Mexico, biogeography); Castro-Aguirre & Balart, 1997: 144 (listed, La Paz Bay, Baja California Sur, Mexico); Bussing 1998: 252–253, 257–259, figures and map 24 (identification key, species account, distribution in Costa Rica); Castro-Aguirre et al., 1999: 234 (synonymy list, distribution in Mexico); Umaña in Lobo & Bolaños, 2005:76 (listed in Costa Rica); Miller et al., 2006:148 (listed, short diagnosis, distribution in Mexico); Vega et al., 2006: 92 (listed in Panamá); Matamoros et al., 2009:16, 35 (in part, listed in Honduras); Angulo et al., 2013: 998 (listed, distribution in Costa Rica).

Pseudophallus starksi starksi: Lüling, 1974: 201 (as a geographic race of P. starksi ).

Pseudophallus lineatus : (non Kaup): Alvarez del Villar, 1970: 80 (new combination, incorrect identification, characters and distribution in Mexico); Dawson, 1982: 47 (synonymic list of P. starksii ).

Diagnosis. Pseudophallus starksii differs from P. brasiliensis , P. mindii and P. galadrielae in having 95.3% of specimens with 14 trunk rings (vs. most usually with 13 trunk rings).

Pseudophallus starksii can be distinguished from P. elcapitanensis by the following combination of characters: 36–44 dorsal-fin rays (89.5% of specimens with 38–41), 49–53 body rings (91.9% of specimens with 50–51), 19–21 brood-pouch rings (64.9% of adult male specimens with 21) and 8.5–10.5 subdorsal tail rings (86.7% of specimens with 9.25–10.0) vs. 30–35 dorsal-fin rays (82.1% of specimens with 31–33), 47–51 body rings (89.8% of specimens with 48–49), 17–19 brood-pouch rings (90.9% of adult males specimens with 17–18) and 6.75–7.5 subdorsal tail rings (97.4% of specimens with 6.75–7.25) in P. elcapitanensis .

Description. Meristic and morphometric characters detailed in Table 2. General appearance and coloration in Figs. 2D and 3D . Body without scales, covered with bony plates forming rings along its axis, bony plates horizontally oval, with an elevation in midline forming bony ridges along body. Trunk rings 13–15 (95.9% of specimens with 14). Tail rings 35–39 (91.1% of specimens with 36–37). Body rings 49–53 (91.9% of specimens with 50–51). Subdorsal tail rings 8.5–10.5 (87.8% of specimens with 9.25–10.0). Subdorsal trunk rings from 0–0.5. Subdorsal tail rings 8.5–10.5 (87.0% of specimens with 9.25–10.0). Dorsal-fin origin at vertical between posterior half of posteriormost trunk ring and anteriormost caudal ring. Posterior tip of superior trunk ridge at vertical through superior tail ridge, between 6.0 and 7.5 anteriormost tail rings. Lateral trunk ridge continuous with superior tail ridge with origin on first caudal ring. Inferior trunk ridge continuous with inferior tail ridge.

Body slender and cylindrical, transversal section of trunk heptagonal, with transition to quadrangular in tail at end of superior trunk ridge. Maximum body depth of females and juveniles at middle of trunk; in males at anterior region of the brood pouch. Body depth decreasing posteriorly along superior trunk ridge to minimum depth on caudal peduncle. Dorsal profile of head straight or slightly inclined at snout region, more inclined at orbital region and straight at posterior region, with two nuchal plates between occipital region and first trunk ring. Ventral profile of head straight.

Dorsal fin not elevated, curved on first rays, straight along median portion with pointed end. Dorsal-fin rays of similar length, except the first two rays. Dorsal-fin rays 36–44 (89.5% of specimens with 38–41), posteriormost two rays closer to each other, difficult to distinguish in some specimens. Pectoral fin small, middle rays longer than outer and inner rays. Pectoral-fin origin on anteriormost trunk ring, and distal margin rarely extending beyond third trunk ring. Pectoral-fin rays 11–15 (68.5% of specimens with 14). Caudal fin small and fan like, with 10 rays (5 rays in each hypural plate). Pelvic and anal fins absent.

Mouth terminal, slightly inclined dorsally, jaws edentulous. Lower jaw slightly prognathous. Snout short, tubular, without dorsal or lateral ridges. Dorsal profile of snout with slight inclination following dorsal margin of eyes. Eyes located longitudinally at middle of head. Opercle occupying most of lateral surface of head posterior to eye, opercular opening reduced to pore on posterodorsal margin of opercle. Opercular ridge generally indistinct, except for some specimens with small protuberance on anterior region of opercle, following superior margin of bone. Lateral line and lateral line canals absent.

Sexual dimorphism. Males with brood pouch. Origin of brood pouch on posteriormost trunk rings, brood pouch rings extending along 19–22 rings (64.8% of specimens 21 brood pouch rings). Bony plates of the brood pouch well developed and T-shaped encasing entire lateral portion of pouch, decreasing in size posteriorly and parallel with the inferior tail ridge. Ventral region of pouch with contralateral dermal folds extending towards the midline, dividing brood pouch longitudinally. Adult females with anal papilla on anal ring with small, barely visible spicules.

Coloration in alcohol. Ground color of head and body light beige to brown, sometimes with horizontal stripe more evident on the head and anterior portion of the body formed by different tones of brown. Smaller stripes on dorsolateral part of head and dorsal part of opercle. Thicker bony regions at body ridges with less or no chromatophores, forming nearly white longitudinal lines. Some specimens with series of darker arch-shaped blotches in lateral view, at every two or four rings. Bony rings with one or two white spots underneath bony plates; shape of spot more circular when single spot present and oval shaped due to proximity of the spots when two spots present. Dorsolateral part of snout with layer of skin and mesethmoid forming light stripe. Dorsal-fin rays with scattered brown chromatophores. Pectoral fin hyaline. Caudal fin with same general coloration of body, sometimes in slightly different tone. ( Figs. 2D and 3D ).

Distribution. Freshwater coastal drainages of the Pacific, from Río Santa Rosa, Ecuador, to San Jose del Cabo, Baja California, Mexico ( Fig. 5 ).

Remarks. A holotype was not designated in the original description of Siphostoma starksii , but Jordan (1895: 381) listed as 2686 the catalog number of the type specimens of the species at the Museum of Stanford University (SU, currently at CAS). Three specimens from this original lot were sent to three different institutions: the United States National Museum of Natural History (USNM 47425), Zoologisches Museum Berlin (ZMB Pisces 14058) and the British Museum, London (BMNH 1895.5.27.321), with five specimens remaining as SU 2686 (now CAS 102686). Later, one additional specimen from lot SU 2686 was dismembered into CAS-SU 67185 (now CAS 167185). Therefore, those eight specimens are part of the original type series (i.e., syntypes) of the species (ICZN, 1999: Art. 72.4.1.1 and 73.2).

In the first part of a series of four volumes of “The fishes of North and Middle America”, Jordan & Evermann (1896a: 771) indicated the lot USNM 47425 as “ type ” of Siphostoma starksi (sic). On the third part of the work by Jordan & Evermann (1898: 2838), SU 2686 is indicated as “ type ” of Siphostoma starksii and in the fourth part of the work by Jordan & Evermann (1900: 3259, pl. CXX) one specimen from lot SU 2686 is illustrated.

Dawson (1982: 45, 49) listed lot SU 67185 (now CAS 167185) as lectotype of Siphostoma starksii . Ferraris (2003: 591) indicated this specimen as being the same of that illustrated in Jordan & Evermann (1900: 3259, pl. CXX) and therefore, credited the lectotype designation of the species to the latter authors. However, the previous mention of the specimen USNM 47425 as type of S. starksii , by Jordan & Evermann (1896a: 771) should be considered as the lectotype designation of the species (ICZN, 1999: Art. 74.5) and other lectotype designations (e.g., Jordan & Evermann, 1900 or Dawson, 1982) are deemed invalid (ICZN 1999: Art. 74.1.1).

Type material: Lectotype. USNM 47425 , 1 , Río Presidio , about a mile below village of Presidio, Sinaloa, Mexico, D. S. Jordan et al., 24 Dec 1894 – 25 Jan 1895 (not examined) . Paralectotypes. CAS 102686 , 4 , 51.0– 123.3 mm SL , CAS 167185 , 1, 132.2 mm SL, same data of lectotype; ZMB 14058, 1, BMNH 1895.5.27.321, 1, same data of lectotype (not examined) .

Non type material. Ecuador: El Oro: CAS 109312 , 1, 117.2 mm SL, River at Santa Rosa, P. O. Simmons 1898-1899 .

Colombia: Valle del Cauca: AMNH 32971 2, 117.3– 124.4 mm SL, quebrada Guangui, 1/ 2 km above Río Patia, upper Río Saija drainage, J. W. Daly, 12 Feb 1973 .

Panama: Darien: MCZ 52992, 1, 64.0 mm SL, Boca de Río Capeti, in fast riffle about 10m upstream of mouth at Río Tuira , 08º05`N 77º35`W, K. E. Hartel, 27 Mar 1978; MCZ 88624, 7 , 112.4 – 146.5 mm SL, Río Picuro, just above confluence with Río Tuira , 08º00`N 77º32`W, W. C. Starnes et al., 16 Feb 1985; USNM 208365 , 6 , 134.9 – 171.1 mm SL, upper tributary of Río Jaque , H. G. Loftin & Dean, 11 Feb 1968; USNM 293603 , 9 , 68.9–135.6 mm SL, Río San Antonio at Garachini , trib. Golfo de San Miguel, J. Lundberg, R. Bouchard & J. Louton, 07 Mar 1988 ;

Panama: UF 12983, 1, 127.8 mm SL, Creek 11 mi. S of Chepo on road to El Llano, Brockman & Child, 29 Apr 1965; USNM 293475 , 8 , 72.3–100.1 mm SL, Pan American Highway ca. 17 Km. E of el Chepo (Bayano Drive), Río Terable , W. Starnes et al., 25 Feb 1985 ; Veraguas: ANSP 104211, 3, 106.0– 124.5 mm SL, Creek Quebrada Piedra at bridge about 8 mi. W. of El Maria on Sona-Remedios road, H. G. Loftin, E. L. Tyson & M. Canipe, 11 Nov 1961; ANSP 104408 , 4 , 63.3 – 116.3 SL, creek at bridge 3 mi. E of Sona-Santiago road, H. G. Loftin & E. L. Tyson 28 Oct 1961 ; Chiriqui: ANSP 104181, 1, 121.9 mm SL, creek into Río Jacaque near San Lorenzo at farm of Dr. Alvarez, H. G. Loftin & E. L. Tyson, 9 Dec 1961; ANSP 119934 , 2 , 54.4–81.6 mm SL, Creek 5 mi. W of Boca del Monte on old road to David, H. G. Loftin & E. L. Tyson 18 Nov 1961; ANSP 119935 , 2 , 63.7–91.1 mm SL, creek at bridge 5 mi W. of San Felix bridge on Interamerican Highway new section, H. G. Loftin & E. L. Tyson 18 Nov 1961; ANSP 11936 , 18 , 50.6–124.7 mm SL, Río Tinta , creek at bridge 3 mi. W of Río Tabasara on Sona-Remedios road, H. G. Loftin, E. L. Tyson & M. Canipe, 11 Nov 1961 : ANSP 146754, 1, 98.8 mm SL, Río Las Vueltas at Interamerican Hwy, st. 51, D. Fromm & P. Fromm, 4 Mar 1981 .

Costa Rica: Puntarenas: FMNH 7901, 1, 83.4 mm SL, Jesus Maria River, S. E. Meek, 1912; FMNH 76970, 1, 98.1 mm SL, Río Grande Terraba , R. K. Johnson, 7 Jul 1973; ROM 88191 , 3, 81.3–138.0 mm SL, Quebrada Benjamin, 1 km West of Palmar Norte, Osa Peninsula , 08º57`54’’N 83º27`44”W, B. Wisenden, 1 May 1991 ; TNHC 15359, 9, 91.6–130.3 mm SL, Río Pavo, upstream from Los Patos trail, P. N. Corcovado , 08º30`29’’N 83º36`03’’W, K. Winemiller, 13 Jul 1983 ; UF 19558, 1, 125.1 mm SL, Río Chires, ca 2 km N off Estarillos , ca 25 air mi. WNW of Quepos, C. Gilbert et al., 2 Jul 1973 ; San José: ANSP 164263, 3,76.1–138.0 mm SL, Río Barú ca 7 km above Dominical, D. Fromm, 11 Mar 1989 ; Guanacaste: UF 19569, 1, 158.8 mm SL, Río Corobia , 5 km NW of Las Canas, near Pan American Hwy, at La Pacifica motel, Gilbert, McAllister & Balon, 4 Jul 1973 .

Honduras: Valle: LACM 32295–11, 1, 135.8 mm SL, Río Nacaome at Pan American Highway, Martin & Brame, 8 Feb 1969 .

El Salvador: La Union: TU 202696, 7, 88.2–131.6 mm SL, trib. to Gulfo de Fonseca , 35.2 miles E of San Miguel, Pan American Hwy, R. D. Sutkus & Smalley, 2 Jun 1962 .

Guatemala: Santa Rosa: AMNH 25100, 1, 87.3 mm SL, Río Margarita, under bridge of Carretera Internacional Del Pacifico , D.E. Rosen, K. Kallman & R. Dorion, 24 Feb 1963 .

Mexico: Oaxaca: FMNH 4717, 1, 104.3 mm SL, San Geronimo, S. E. Meek, 29 Feb 1903 ; Guerrero: UMMZ 178462, 8, 69.2–103.0 mm SL, Río Coyaquilla , 60 road miles SE of Zihuatenejo, R. R. Miller & M. Miller, 21 Mar 1957 ; Nayarit: UMMZ 172107, 8, 87.3–120.1 mm SL, Río La Lima at La Lima , 7.5 mi by road SSW of Las Varas, R. R. Miller & J. T. Greenbank, 24 Feb 1955 ; UMMZ 178588, 10, 85.0– 100.4 mm SL, Río Rosario at Hwy. 15 bridge near Rosario , Pacific Dr, R. R. Miller & M. Miller, 08 Apr 1957 ; Baja California Sur: CAS ( SU) 5943, 4, 67.1–124.9 mm SL, San Jose del Cabo, Eisen & Vaslit .

Figures

FIGURE 2: Species of the genus Pseudophallus: A—Pseudophallus brasiliensis (paratype) USNM 212058, 69.1 mm SL, Igarapé Inó, Furo de Panaquera, Rio Tocantins, Pará, Brazil. B—Pseudophallus elcapitanensis MCZ 88623, 110.6 mm SL, Río Picuro, Darien, Panama. C—Pseudophallus mindii (holotype) USNM 81770, 90.3 mm SL, Mindi Creek, Canal Zone, Panama. D—Pseudophallus starksii USNM 208365, 171.1 mm SL, upper tributary of Río Jaque, Província de Darien, Panama. E—Pseudophallus galadrielae. (holotype) FMNH 126156, 63.1mm SL, Lago Izabal, Guatemala. Red arrow indicates the anal papilla.

FIGURE 3: Detail of anterior portion of body of species of the genus Pseudophallus A—Pseudophallus brasiliensis (paratype) USNM 212058, 69.1 mm SL, Igarapé Inó Furo de Panaquera, Rio Tocantins, Pará, Brazil. B—Pseudophallus elcapitanensis MCZ 88623, 110.6 mm SL, Río Picuro, Darien, Panama. C—Pseudophallus mindii (holotype) USNM 81770, 90.3 mm SL, Mindi Creek, Canal Zone, Panama. D—Pseudophallus starksii USNM 208365, 171.1 mm SL, upper tributary of Río Jaque, Província de Darien, Panama. E—Pseudophallus galadrielae. (holotype), FMNH 126156, 63.1 mm SL, Lago Izabal, Guatemala.

FIGURE 5. Distribution map of species of Pseudophallus occurring in drainages along the Pacific coast based on specimens examined in this study.

FIGURE 7. Maps of distribution of specimens of Pseudophallus available for study. Left side: distribution of specimens examined for the morphological analysis in this study. Right side map: distribution of voucher specimens with genetic sequences on GenBank (as of April 2020).